Geo log i cal Quar terly, 2020, 64 (4): 876–897 DOI: http://dx.doi.org/10.7306/gq.1565

The his tory of (Ursidae, , Mammalia) from Silesia (south ern Po land) and the neigh bour ing ar eas

Adrian MARCISZAK1, * and Grzegorz LIPECKI2

1 Uni ver sity of Wroc³aw, De part ment of Palaeozoology, In sti tute of En vi ron men tal Bi ol ogy, Fac ulty of Bi o log i cal Sci ences, Sienkiewicza 21, 50-335 Wroc³aw, Po land 2 Pol ish Academy of Sciences, Insti tute of System atics and Evolu tion of An i mals, S³awkowska 17, 31-016 Kraków, Po land

Marciszak, A., Lipecki, G., 2020. The history of bears (Ursidae, Carnivora, Mammalia) from Silesia (south ern Poland) and the neigh bour ing ar eas. Geo log i cal Quar terly, 64 (4): 876–897, doi: 10.7306/gq.1565

As so ci ate Ed i tor: Micha³ Zatoñ

Revi sion of the Silesian fauna, based on ma terial from 152 sites, mainly cave and karstic local i ties, and also ar chae o- log i cal and open-air sites, shows the presence of 13 forms and spe cies. These re cords en com pass the last 16.5 Ma and may be di vided into five main morphophyletic groups. The old est bears, rep resented by the genera Ballusia and , are dated to 16.5–11 Ma, and belonged to the stem forms of the subfamily . Af ter a break of 6 My, the earli est mem bers of the genus appeared, and all known Plio cene bears in Silesia be long to the ge nus Ursus were identi fied as . They repre sent one or two migra tion events. Ad di tion ally, between 3.6 and 3.2 Ma, a single oc cur rence of insigne was recorded from the Wê¿e 1 site. A few Early Pleis tocene bear records are repre sented by U. etruscus, which was a proba ble an ces tor of both arctoid and spelaeoid bear lin eages. The old est rep resen tative of Ursus ex gr. arctos known so far and assigned to U. a. suessenbornensis is known from one lat est Early Pleis tocene (1.2–0.9 Ma) lo cal ity, while other Pol ish re cords of this form re quire con firma tion. The first oc cur rence of U. deningeri, the old est taxon within the U. ex gr. deningeri–spelaeus lin eage, was recorded from ~700 ka de pos its in Silesia. Dur ing the pro nounced cold period of MIS 12, the Scan di navian ice sheet cov ered al most the en tire mod ern terri tory of Po land, with the excep tion of the Sudetes and the Carpathians. The ac com pa nying dras tic fau nal turn over led to the forma tion of the pan-Eurasian Mam moth Fauna at ~460 ka. At that time a char ac teris tic mem ber of this fauna, the steppe Ursus arctos priscus, a spe cific ecomorph adapted to live in open grass lands, ap peared in this re gion. It sur vived un til the be gin ning of MIS 1, when mod ern Ursus arctos arctos ap peared in Silesia and sur vived to the pres ent day. U. deningeri was the most com mon bear during the Mid dle Pleisto cene, while the first re cords of U. spelaeus spelaeus ap peared since MIS 7. The lat ter form was re placed by U. ingressus during the Late Pleisto cene (~110–100 ka). Spelaeoid bears to tally domi nated the cave as sem blage, and fi nally van ished be tween 27 and 24 ka.

Key words: taxon omy, Miocene, Pliocene, Pleisto cene, ursid lineages.

INTRODUCTION Bears spread all over the North ern Hemi sphere dur ing the Mio cene and gave rise to a con sid er able num ber of spe cies, from which a single one has sur vived to the pres ent in Po land. Finds of dif fer ent rep re sen ta tives of the family Ursidae are The Silesian his tory of bears started with the earli est Pol ish re - fre quently made in Po land, and their high est den sity has been cord of Ursavus elmensis from the Przeworno 2 site dated to re corded from the cave and karstic sites of Silesia and im me di - 16–14.5 Ma. This taxon is also gen er ally consid ered to be the ately neighbour ing ar eas. Palaeontologically this re gion is one ear li est un dis puted bear spe cies (Baryshnikov and Lavrov, of the most pro lific in Po land. Bear re cords from Silesia and 2015). The Mio cene ge nus Ursavus Schlosser, 1899 is re - nearby areas al low us to re con struct the re gional his tory of the garded as a di rect an ces tor of mod ern ur sine bears of the ge - fam ily over the last ~16 My. This timespan is, though, in com - nus Ursus Linnaeus, 1758. Bears in Silesia have been al most pletely cov ered, and there are some gaps, es pecially in the mid - ex clu sively rep re sented by spe cies of Ursus over the last 4 My, dle Mio cene to Mid dle Pleis to cene in ter val. with the ex cep tion of a sin gle oc cur rence of Agriotherium that per sisted from the Mio cene (Stach, 1953, 1957; Wolsan, 1989). Ur sine bears have existed in Silesia con tin u ously since the Mid dle Plio cene, from ~4 Ma. Al though only sporadic in some in ter vals, bears be came one of the dom i nant mam mal groups * Cor re spond ing au thor, e-mail: [email protected] dur ing the Mid dle and Late Pleis to cene. This pa per pro vides a Received: April 14, 2020; ac cepted: Sep tem ber 17, 2020; first gen eral over view of the his tory and taxon omy of ur sine bears in pub lished on line: November 12, 2020 Silesia and the near est neigh bour ing ar eas, as a crit i cal re view The his tory of bears (Ursidae, Carnivora, Mammalia) from Silesia (south ern Poland) and the neigh bouring ar eas 877 of the lit er a ture com ple mented by our own re search re sults. fam ily in the palaeofaunas grew sig nif i cantly, and from the Mid - Size and mor pho log i cal anal y ses are sup ple mented by ura - dle Mio cene bears be came im por tant mem bers of the nium-tho rium (UTh) and ac cel er a tor mass spec trom e try (AMS) palaeoguilds. By that time there had al ready been estab lished a dat ing meth ods as well as by DNA analy sis. model of bear evo lu tion, based on a large, solid an i mal, mostly om niv orous or car niv orous, able to com pete with even the larg - est ac tive carni vores like big cats. In the last 15 My on almost all MATERIAL AND METHODS con ti nents (ex pect Aus tra lia) a large bear, Agriotherium ear lier, Arctodus or Ursus later, which played an impor tant role as a The sub di vi sion of the fam ily Ursidae Fischer, 1814 into five scav en ger of and com pet i tor to other large car ni vores. subfamilies: Ursinae Fischer, 1814, Arctotheriinae F. Dur ing this evo lu tion, the size of mem bers of the Ursinae Ameghino, 1903, Grevé, 1892, Agriotheriinae subfamily also grew con sid er ably (Fig. 2). The ear li est Silesian Kretzoi, 1929, and Ursavinae Kretzoi, 1945 is made sensu bear, B. elmenensis, was the size of a lynx, while their di rect de - Wag ner (2010). Ac cord ing to this con cep tion, all Eur asian rep - scen dants of the ge nus Ursavus were al ready the size of a re sen ta tives of the Ursinae subfamily discussed in this pa per large wolf. Plio cene forms like U. minimus were com pa ra ble in may be assigned to the genus Ursus. The tax onomic overview size with the modern Hi ma la yan black bear Ursus thibetanus was made ac cord ing to main groups of Eur asian bears sensu Cuvier, 1823. De spite its rel a tively large di men sions, A. in signe Mazza and Rustioni (1994): (1) minimus–thibetanus, (2) from the Wê¿e 1 site is a me dium-sized mem ber of the genus etruscus, (3) arctos, and (4) deningeri–spelaeus. The def i ni tion Agriotherium. The Early Pleis to cene U. etruscus reached the and sub di vi sions of the Qua ter nary Pe riod fol low Gibbard and size of a modern Cen tral Eu ro pean brown bear. Ursids reached Co hen (2008) and Gibbard and Head (2009a, b). The def i ni tion their maxi mum size in the Mid dle and es pe cially Late Pleis to - and sub di vi sions of mam mal zones and their cor re la tion with cene, where deningeroid and spelaeoid bears dom i nated in the chronostratigraphic scale as well as the MN-zones fol low caves and were the larg est Eur asian bears. Com pa ra ble in size Gliozzi et al. (1997) and Fejfar et al. (1998). The no men cla tur al with them, and even ex ceed ing them as regards large males, cod i fi ca tion fol lows the 4th edi tion of the In ter national Code of was the larg est car ni vore in Silesia and in deed in Eur asia in Zoo log i cal No men cla ture (ICZN, 1999). Cap i tal and low er case gen eral, the Ice Age steppe brown bear U. a. priscus (Fig. 2). let ters, P/p (premol ars), and M/m (mo lars), re fer to upper and lower cheek-teeth, re spec tively. The area in cluded in the anal ysis co in cides with the histor i- MIOCENE (16–5 Ma) cal bound aries of Silesia (Provinz Schlesien) as shown by Pax (1921, 1925), with some mi nor mod i fi ca tions (Fig. 1). At pres- The old est un dis puted Silesian bear spe cies is Ballusia ent, Silesia cov ers ~50,000 km2; it ex tends along the Odra River elmensis (Stehlin, 1917) from the Przeworno 2 site dated to and is mostly located in Po land, with small parts in the Czech 16–14.5 Ma. This form oc cu pied a stem po si tion among ur sine Re pub lic and Ger many (Fig. 1; Czapliñski et al., 2007). taxa (Qiu et al., 2014). The genus ap par ently dispersed from Ra dio car bon dates were made in the Poznañ Ra dio car bon Asia into North Amer ica and Eu rope at ~20 Ma, as shown by a Lab o ra tory fol low ing their pre-treat ment pro to col for the ex trac - large num ber of fos sils in Eu rope grad ing to ward east ern Asia tion of col la gen (Longin, 1971; Piotrowska and Goslar, 2002). (Qiu et al., 2014). This small bear (com para ble in size to the Be fore ex trac tion, the de gree of col la gen deg ra da tion is Eur asian lynx Lynx lynx (Linnaeus, 1758) is char ac ter ized by checked by mea sur ing the con tent of ni tro gen and car bon in pro por tion ally small M2 with short and nar row talon. Ac cord ing bone, us ing a Flash EA 1112 Se ries (Thermo-Sci en tific) to Baryshnikov and Lavrov (2015), B. elmensis was prob a bly a analyser. The sam ples are re garded suit able for col la gen dat - small ar boreal an i mal with an om niv orous diet. This spe cies is ing if ni tro gen con tent in bone is ³0.6%, and the C/N ra tion is £5. rarely noted in Cen tral Eu rope, for in stance at the Ger man sites Suit able bones are crushed me chan i cally to gran u la - Wintershof-West (type local ity for MN3; Dehm, 1950) and tion<0.3 mm, the bone pow der is then treated with 2M HCl Hambach 6C (Mörs et al., 2000) (room temp., 20 min), and 0.1M NaOH (room temp., 1 h). After True mem bers of the genus Ursavus, U. brevirhinus each step of treatment, the sam ple is cen tri fuged and the resid - (Hofmann, 1887) and U. primaevus (Gaillard, 1899), are the uum is col lected. Extrac tion of col la gen is pro cessed in HCl next two ur sine bears re corded from Silesia. Both spe cies can (pH = 3.80°C, 10 h), and af ter cen tri fug ing the re sid uum is re - be re garded as the de scen dants of B. elmensis (sensu moved. The extracted col la gen is then ultra fil tered on Ginsburg and Morales, 1998), and were re corded mostly from pre-cleaned Vivaspin 15 MWCO 30 kD fil ters. The qual ity of the cen tral Silesia. U. brevirhinus had a long timespan from col la gen is ul ti mately as sessed based on the C/N atomic ra tio 18–11 Ma (MN 4-9). This spe cies was widely distrib uted across (in ter val of ac cep tance: 2.7–3.5) and col la gen ex trac tion yield Eu rope: Steieregg and Voitsberg (MN 5), and Göriach (MN 6), (ac cep tance thresh old: 0.5%). On de mand, carbon and ni tro - all in Aus tria (Hoffmann, 1887; Koken, 1888; Hoffmann, 1892; gen sta ble iso to pic com po si tion of the col la gen can be de ter - Schlosser, 1899a, 1899b; Thenius, 1949a, 1949b); Opole 1 mined (Bronk Ramsey et al., 2004). (14–13 Ma) and Gliwice-Soœnicowice in Po land (14–11 Ma) (Wegner, 1908, 1913; Koenigswald, 1925; Helbing, 1936), and Can Llobasteres (11.1–9.7 Ma) in Spain (Crusafont-Pairo and TAXONOMIC PART Kurtén, 1976). U. brevirhinus from Can Llobasteres rep re sents the larg est and the most ad vanced spec i men, while in di vid u als Re vi sion of the Silesian and neigh bour hood bear fauna, from Gliwice-Soœnicowice strongly varied in size. Ac cord ing to based on 152 sites, mainly cave and karstic, but also ar chae o - Qiu et al. (2014), they appar ently com prise two forms, small log i cal and open-air, showed the pres ence of 13 bear taxa and large, and ac cord ing to them only the small maxilla with (Fig. 1 and Appen dix 1*). The re cords covered the last 16 Ma, M1-M2 in Wegner (1913: pl. XII-16) can be assigned to U. and were di vided into five main groups. The im por tance of this brevirhinus. This in ter pre ta tion can not be en tirely ruled out,

* Supplementary data associated with this article can be found, in the online version, at doi: 10.7306/gq.1565 878 Adrian Marciszak and Grzegorz Lipecki

Fig. 1. Local i ties in Silesia and neighbour ing areas with bear remains, discussed in the text, in chrono log i cal order The Silesia bound ary is su perim posed on mod ern national bor ders, in di cated by black lines. State bound aries are in di cated by red lines, and admin is trative bound aries are in di cated by blue lines. In the in sert showing the lo ca tion of Silesia in Eu rope, the Silesia bound ary (in red) is su per im posed on mod ern na tional bor ders; Mio cene: 1 – Przeworno 2; 2 – Be³chatów B; 3 – Opole 1; 4 – Opole 2; 5 – Gliwice–Soœnicowice; Plio cene: 6 – Pañska Góra; 7 – Wê¿e 1; 8 – Wê¿e 2; 9 – Rêbielice Królewskie–Kamienio³om, by the cemetary; Early and Middle Pleisto - cene: 10 – Jasna Strzegowska cave; 11 – ¯abia cave; 12 – Zamkowa Dolna cave, fauna C; 13 – Tunel Wielki cave; 14 – Przymi³owice C; 15 – Po³udniowa cave; 16 – Draby 2, 3, 5, 8 and 9; 17 – Biœnik cave, layers 19ad-14; 18 – Deszczowa cave, layers 1–4; 19 – Nietoperzowa cave, lay ers 18–14; 20 – Stajnia cave, layers H–G; 21 – Komarowa cave, lower most layers; 22 – Ciemna cave, layers 19–17; Late Pleis to cene: 17 – Biœnik cave, layers 12–1; 18 – Deszczowa cave, layers 5–8; 19 – Nietoperzowa cave, layers 13–3; 20 – Stajnia cave, layers F–B; 21 – Komarowa cave; 22 – Ciemna cave, layers 16–2; 23 – Dziadowa ska³a cave, layers 3–4; 24 – Imbramowice; 25 – Koziarnia cave; 26 – NiedŸwiedzia cave; 27 – Miniaturka cave; 29 – Wschodnia cave; 30 – Naciekowa cave; 31 – Pó³nocna Du¿a cave; 32 – by Wschodniej cave; 33 – caves of Mt Po³om; 34 – Wierzchowska Górna cave; 35 – Wylotne shelter; 36 – NiedŸwiedzia Górna cave; 37 – £okietka cave; 38 – Perspektywiczna cave; 39 – Winna Góra; 40 – Skarszyn; 41 – Towarna cave; 42 – Mamutowa cave; 43 – Pyskowice; 44 – Rakoszyce; 45 – Paw³owiczki; 46 – Solna Jama cave; 47 – cave 4 on Mt Birów; 48 – Bêb³owska Dolna cave; 49 – caves near Olkusz; 50 – cave near Horní Lánov; 51 – Schöpstal–Kunnersdorf; 52 – cave 5 in Z³oty Potok; 53 – Grota NiedŸwiedzia cave; 54 – Dzier¿no; 55 – G³ówna w Kopcowej Górze cave; 56 – Gorenicka cave; 57 – K¹ty Wroc³awskie; 58 – Koralowa cave; 59 – Lubliniec; 60 – £abajowa cave; 61 – Murek cave; 62 – na Go³¹bcu cave; 63 – nad Galosk¹ cave; 64 – na Gaiku 2 shel ter; 65 – na Mi³aszówce cave; 66 – na Wrzosach Pó³nocna cave; 67 – nad Matk¹ Bosk¹ cave; 68 – Okiennik cave; 69 – pod Kochank¹ cave; 70 – Poœrednie shel ter; 71 – Rogó¿ka cave; 72 – Sadlana cave; 73 – shel ter no 1 in Srocko; 74 – Wiêksze w Krzy¿owej Skale; 75 – w Ogrójcu Górne shel ter; 76 – w Okopach Wielka Dolna cave; 77 – shel ter no. 6 in Z³oty Potok; 78 – shel ter no. 7 in Z³oty Potok; 79 – Sypialnia cave; 80 – Wiercica cave; 81 – Zbójecka cave; 82 – Zegar cave; 83 – Ziêbice; 84 – Zittau (¯ytawa);85 – Kraków Spadzista; 86 – Maszycka cave; Ho lo cene: 15 – Po³udniowa cave; 17 – Biœnik cave, layer 1ab; 18 – Deszczowa cave, layers 9–11; 19 – Nietoperzowa cave, layers 2–1; 20 – Stajnia cave, layers B–A; 21 – Komarowa cave, up permost layers; 22 – Ciemna cave, layers 2–1; 23 – Dziadowa ska³a cave, layers 9–13; 26 – NiedŸwiedzia cave; 29 – Wschodnia cave; 30 – Naciekowa cave; 32 – by Wschodniej cave; 87 – Aven in Po³omie cave; 88 – Babie Nogi shel ter; 89 – Berkowa cave; 90 – Cisowe 1 shel ter; 91 – Cisowe 2 shel ter; 92 – Panna shel ter; 93 – Trwoga Paleontologa shel ter; 94 – G³uszyca–Górna; 95 – Górne shel ter above Stajnia cave; 96 – Krucza Ska³a shel ter; 97 – Maurycego cave; 98 – Kryszta³owa cave; 99 – Medvìdia cave; 100 – Ruska Ska³a shel ter; 101 – Olsztyñska and Wszystkich Œwiêtych caves; 102 – Studnisko cave; 103 – Trzebniowska cave; 104 – Wierna cave; 105 – Wilczy Dól shel ter; 106 – Œwiêtej Anny Mt.; 107 – Œlê¿a Mt; 108 – Szkaradowo; 109 – G³ogów–Nosocice; 110 – Miechów; 111 – Wilcze shel ter no 3; 112 – Domas³aw; 113 – Kamieñ Œl¹ski; 114 – Rogowiec casttle; 115 – Borsuka cave, layer 3; 116 – cave no. 3 on Mt Birów; 117 – Grodziec; 118 – £azy; 119 – Kietrz; 120 – on Wrzosach Po³udniowa cave; 121 – Grzybiany 1; 122 – Dzia³oszyn; 123 – Ojców cas tle; 124 – Karczna Góra; 125 – shel ter no. 3 in Smoleñ; 126 – in Kostkowice shel ter; 127 – Zielona cave; 128 – Góra 1; 129 – Wroc³aw Marii Magdaleny street; 130 –in Zielonej Górze cave; 131 – Strachów; 132 – Opole; 133 – Trzebnica; 134 – Rzymówka; 135 – Kraków–Square Market; 136 – Legnica; 137 – Dêbnica; 138 – Racibórz; 139 – Santok; 140 – Szczerba cas tle; 141 – Sieradz; 142 – Kaszowo 1; 143 – Krosno Odrzañskie; 144 – Niemcza; 145 – Kêdzierzyn-Ko¿le; 146 – Wroc³aw–Ostrów Tumski; 147 – Bytom Odrzañski; 148 – Opole–Ostrówek; 149 – Miêdzyrzecz 1; 150 – Grodziszcze; 151 – Wroc³aw–Nowy Targ; 152 – Ma³gorzaty shelter The his tory of bears (Ursidae, Carnivora, Mammalia) from Silesia (south ern Po land) and the neigh bouring ar eas 879

Fig. 2. Size compar i son of dif ferent Silesian bears Ballusia elmensis – Przeworno 2 (16–14.5 Ma), Ursavus brevirhinus – Opole 1 (14–13 Ma), Ursus minimus and Agriotherium insigne – Wê¿e 1 (3.6–3.2 Ma), U. etruscus – ¯abia cave (1.7–1.5 Ma), U. arctos arctos – Wschodnia cave (14.3–13.2 ka), U. deningeri – Draby 3 (440–400 ka), U. rossicus – Naciekowa cave (60–40 ka), U. spelaeus spelaeus – Biœnik cave, layers 15–13 (200–130 ka), U. ingressus and U. arctos priscus – NiedŸwiedzia cave (100–35 ka); all bears are adult males shown at the same scale, com pared with a man 1.8 m high though an al ter na tive ex pla na tion of sex ual di mor phism or dents, 14 car ni vores, 2 perissodactyls and 2 ar tio dac tyls) intraspecific vari abil ity is also pos si ble. This spe cies has also (Samsonowicz, 1934; Stach, 1953, 1957, 1959; Nadachowski been noted in some far un des cribed ma te rial from Be³chatów B et al., 1989; Kowalski, 1989a, b; Wolsan, 1989; Kowalski, 1990; (14.5–13.8 Ma, Kowalski and Rzebik-Kowalska, 2002), sit u ated Rzebik-Kowalska, 2009; Stefaniak, 2015), and re mains of two ~50 km north of the his tor i cal bound ary of Silesia dif fer ent bears have been found. Among these, bro ken up per The sec ond spe cies, U. primaevus ap peared later in the teeth and better pre served lower i3 and m1 of a big bear-like an- fos sil re cord, at be tween 15 and 11 Ma (MN6-9); it has been i mal have been re corded. Based on a few frag ments (right noted from 4 lo cal i ties within Silesia. Firstly, it was de scribed maxilla with M1-M2, left M2, right P4 and M1, left man di ble, left from Opole 2, sit u ated 1 km from Opole 1, dis cov ered by re- i3 and left and right m1), that prob ably belonged to one in di vid - search ers of the De part ment of Palaeozoology, Uni ver sity of ual, Stach (1957) es tab lished a new, rather poorly defined spe - Wroc³aw. Opole-2 is be lieved to be some what younger cies Agriotherium intermedium. Mem bers of the genus (12–11 Ma) than Opole 1, and some au thors sug gested pos si - Agriotherium were widely distrib uted across Eu rope, In dia, ble co-oc cur rence of U. primaevus and U. brevirhinus (Wolsan, China, North Amer ica and South Af rica, and ex isted be tween 1989; Kowalski, 1990). This is of course pos si ble, but more a 13.5 and 2.5 Ma (Hendey, 1972, 1974; Jiangzuo and Flynn, better expla na tion is that the two spe cies come from two 2019). Af ter the first de scrip tion by Wag ner (1837) as stratigraphically dif fer ent lev els. These spe cies dif fer in a num - Hyaenarctos, and later amended to Agriotherium by Sellards ber of size-re lated and mor pho log i cal fea tures, and over all (1916), with in creas ing finds of new ma te rial, mostly frag men - com par i son of both forms shows the younger evo lu tion ary level tary, Agriotherium grad u ally be came a large het er o ge neous ge - of U. primaevus. When com pared with U. brevirhinus, nus. After Stach’s (1957) pa per, nothing was done with the U. primaevus is larger (m1 length is 15–18 mm for Wê¿e 1 ma te rial, and later au thors (e.g., Wolsan, 1989; U. brevirhinus and 17–20 mm for U. primaevus), P3/p3 and p4 Kowalski, 1990) con tin ued to use the name A. intermedium are broader and have equal crown heights, P4 lacks a with out pay ing at ten tion to the in suf fi cient de scrip tion. Only re - parastyle, the talon of M2 is pro por tion ally broader and longer, cently, af ter a de tailed re vi sion of old, and ad di tion of some and m1 has a shorter and more oblique paraconid, a less open new, ma te rial, it be came clear that this name is no lon ger valid val ley between the paraconid and metaconid, a rel atively higher as a sep a rate spe cies and should be synony mised with protoconid with more steeply mesial and dis tal slopes, and a Agriotherium insigne (Gervais, 1859); this was al ready pointed shorter talonid (de Bonis et al., 2018). Be sides Opole 2, out by Jiangzuo and Flynn (2019). Agriotherium from Wê¿e 1 is U. primaevus in Silesia has also been re corded from slightly smaller than A. in signe from other lo cal i ties, but pre - Gliwice-Soœnicowice, by Qiu et al. (2014) based on large spec i - served dentition shows the same mor pholog i cal fea tures, such mens de scribed pre vi ously by Wegner (1913) as be long ing to as a well-de vel oped cingulum. The small ob served dif fer ences U. brevirhinus. Be sides Silesia (Po land), this spe cies has also be tween Agriotherium from Wê¿e 1 and A. in signe may be ex - been noted from La Grive-Saint-Alban in France (Gaillard, plained due to geo graphic or chro no log i cal dif fer ences, since 1899; Depéret and Llueca, 1928), and Can Ponsich in Spain this site is younger than other Eu ro pean re cords of this spe cies (Crusafont-Pairo and Kurtén, 1976). (Jiangzuo and Flynn, 2019). In some fea tures, Agriotherium from Wê¿e 1 re sem bles also Agriotherium roblesi Mo rales and Aguirre, 1976, known from the Span ish site Venta del Moro PLIOCENE (5–2.58 Ma) (6.8–4.9 Ma, Mo rales and Aguirre, 1976). However, in other mor pho log i cal fea tures, and above all in size, A. roblesi, which Sub se quently in the his tory of Silesian bears there is a wide is a huge bear, so dif fers from Agriotherium from Wê¿e 1 that (6–7 My) gap with out any fos sil ma te rial un til the Late Plio cene can not be as signed to this spe cies. lo cal ity Wê¿e 1, dated to 3.6–3.2 Ma. This re mark able lo cal ity From its great size and some den tal and cra nial fea tures has yielded 115 ver te brate spe cies (in clud ing 9 am phib i ans, 15 like a short broad ros trum, pre-masseteric fossa on the man di - rep tiles, 4 birds, 30 in secti vores, 11 bats, 1 lagomorph, 27 ro - ble and ro bust sec to rial car nassi als, some pre vi ous au thors 880 Adrian Marciszak and Grzegorz Lipecki

(e.g., Kurtén, 1967; Hendey, 1972, 1980) de scribed pro posed by Morlo and Kundrát (2001) and Baryshnikov Agriotherium as highly car niv o rous an i mal pre dat ing large ter - (2007); the latter au thor has named this form as Ursus minimus res trial mam mals. However, other au thors, in view other fea - boeckhi (Schlosser, 1899). This con trasts with the opin ion of tures such as long, gracile limbs, have sug gested this an i mal Wag ner et al. (2011) who sup posed that these bears com pose was highly carniv orous, but obtained most of its diet by scav - a sep arate migra tion event and prob ably also repre sent a sep a - eng ing (Sorkin, 2006; Oldfield et al., 2012). A large rate spe cies. Agriotherium looks like a sav age car ni vore able to catch al most While U. minimus has been re corded only from Perpignan any ani mal in its hab i tat. But this first im pres sion is de cep tive, at ~4.2–3.6 Ma (MN15a) (France, Wag ner, 2010), this spe cies as Agriotherium has a pro por tion ally short but broad snout, typi - be came more nu mer ous and spread across Eu rope from Eng - cal for the car ni vores with strong jaws. Anal ysis has shown that land to the Cau ca sus be tween 3.6 and 3.2 Ma (MN15b). Local i - Agriotherium had one of the stron gest bite forces known among ties with the most rep re sen ta tive ma te rial are Kvabebi the mem bers of the or der Carnivora (Sorkin, 2006). By being (3.2–2.5 Ma; Vekua, 1972), Odesa Cat a combs (3.5–3 Ma, able to crack open bones, this an i mal could ac cess one of the Rosin, 1956), Osztramos 7 (2.8–2.5 Ma; Jánossy, 1978), most nu tri tious parts of an an i mal – the bone mar row – which Gaville (3.2–2.5 Ma; Berzi, 1966), Layna (3.6–3.2 Ma; Soria can last for sev eral years af ter an an i mal’s death when encased and Mo rales, 1976), Les Etouaires (2.8–2.5 Ma; Mazza and in side the bones. However, postcranial bones of Agriotherium Rustioni, 1994) and Wê¿e 1 (Ryziewicz, 1969). Ac cord ing to char ac ter ise it as a large but rel a tively un der pow ered an i mal Wag ner (2010), these bears rep re sent a new (at least a sec - that does not seem to have the skel etal frame work nec es sary to ond) mi gra tion wave of black bears into Eu rope. They were a lit - cope with high stresses, such as those expected to be en coun - tle big ger and more ad vanced in tooth mor phol ogy than their tered while un der go ing ex treme phys i cal ex er tion dur ing catch - ear lier rel atives. Be tween MN 15b to MN 16b, they evolved with ing and sub duing large strug gling prey. Putt ing that all to gether, small changes, and the fos sil ma te rial of U. minimus from Wê¿e it may be con cluded that Agriotherium was the first Silesian 2 (2.4–2.2 Ma) differs from the older bears of Wê¿e 1 mostly in spe cial ised scaven ger. Be ing larger than any other car ni vore be ing slightly larger, while mor pho log i cally these spec i mens are within the fauna from Wê¿e 1, A. in signe could in ef fect have very sim i lar. scared smaller car ni vores away from car casses of their prey, and ob tained this food by kleptoparasitism. This bear may have also sup ple mented its diet with plants and, as one of the first EARLY (~2.0–0.9 Ma) spe cial ised scav en ger bears, was even tu ally re placed by more advanced forms of an other bear ge nus, as well as possi bly by Af ter ~2.0–1.8 Ma U. minimus Devèze & Bouillet, 1827 was other bone crunch ing an i mals such as hye nas. A. in signe has re placed in Eu rope by (Rustioni and Mazza, been re corded from two older, dated to 4.9–4.2 Ma (MN14) lo - 1993b; Wag ner, 2010; Wag ner et al., 2011). The lat est oc cur - cal i ties, i.e. Montpellier in France (Gervais, 1853, 1859a, b; rence of U. minimus in Eu rope is known from the Poggio Rosso Stehlin, 1907; Viret, 1939) and Alcoy-Mina in Spain (Montoya et lo cal ity in It aly (Mazza et al., 2005), as well as from Villány 3 al., 2006). In this con text, the Pol ish re cord from Wê¿e 1 may (Kormos, 1937; Wag ner, 2010; Wag ner et al., 2011) and prob a- be re garded as one of the very late Eu ro pean oc cur rences of bly also Kisláng in Hun gary (Kretzoi, 1954; Wag ner, 2010; this spe cies (and of mem bers of the genus Agriotherium as Wag ner et al., 2011). An Asi atic newcomer, U. etruscus ap- well), fol lowed only by those from the Vialette lo cal ity in France peared in Eu rope from ~2–1.8 Ma and became a con stant pres - (3.2–3.1 Ma, Helbing, 1932; Lacombat et al., 2008). ence at Eu ro pean Early Pleis to cene lo cal i ties. Be cause the Pol - The pro cess of for ma tion of the mod ern Silesian mam mal ish ma te rial of this spe cies is re vis ited in de tail in the sec ond pa- fauna started at ~4.5–4 Ma. It was the time of ap pear ance of the per in this vol ume, here we only note that this spe cies has been first mod ern mam ma lian gen era as well as start of the de vel op - re corded from three Pol ish sites, all lo cated in Silesia. ment of mam ma lian as sem blages sim i lar or iden ti cal to mod ern The in ter val be tween 1.2 and 1.0 Ma in Eu rope was a time ones. Among them, car ni vores were very di verse, and bears when the ear li est rep re sen ta tives of two well-known bear lin - became an in te gral part of these fau nas from the be gin ning. eages, namely arctoid and spelaeoid, ap peared and be came How ever, the exact time and place of the or i gin of this lin eage dom i nant in Eu ro pean fau nas over the next 1 Ma. It was also a re main un known. The Wê¿e 1 lo cal ity has yielded, in ad di tion to time when both lineages started to evolve in de pend ently, lead- A. in signe, also sec ond, a much smaller bear species. Ursus ing to omnivory and oppor tun ism in arctoid bears and herbivory minimus, de scribed orig i nally as Ursus wenzenis (Stach, 1953) and spe ciali sa tion in spelaeoid bears. The ear li est speci mens is better stud ied and de scribed on the ba sis of more abun dant are still very sim i lar, and it is not al ways easy to dis tin guish ma te rial (at least 250 skel etal el ements such as skulls, man di - arctoid and spelaeoid bears. Rustioni and Mazza (1993b) sug - bles, isolated teeth and other bone frag ments; Ryziewicz, gested that mem bers of Ursus ex gr. arctos oc cur al ready 1969). This ear li est rep re sen ta tive of the ge nus Ursus in Eu - around 1.3–1.0 Ma. Lat ter the same ma te rial was re-ex am ined rope first ap peared in the Early Pliocene (4.9–4.7 Ma, MN14) in and as signed to other forms. The bear from Vallonet was identi - the Montpellier lo cal ity in France, from which the mo lars still fied as U. deningeri von Reichenau, 1904 in Baryshnikov bear some Ursavus-like char ac ters (Wag ner, 2006, 2010). This (2007), and from Pirro Nord as U. etruscus Cuvier, 1823 bear is known from a few sites lo cated mostly in SE Eu rope, (Petrucci and Sardela, 2009). All these con cepts were impor - dated to 4.5–4.2 Ma: Kuchurganian beds (Ukraine, Korot - tant breaks in the concepts of Eu ro pean bear tax onomy and kevich, 1967), Baraolt-Cãpeni, known also as Barót-Köpecz phy log eny (Wag ner, 2010). (Ro ma nia, Schlosser, 1899a, b, c; Maier von Mayerfels, 1929), Excel lent ma te rial from Deutsch-Altenburg (most of the and Dorkovo (Bul garia, Delson et al., 2005), but also from Ibe - bones came from DA4B, dated to 1.2–1.1 Ma; oth ers are from rian sites such as Alcoy-Mina (Spain, Montoya et al., 2006), We DA2C1 and DA49, dated to 1.4–1.2 Ma) was regarded as an have also as signed a few iso lated teeth (mostly of limited tax o - early repre sen ta tive of the brown bear Ursus arctos nom i cal value) from Pañska Góra, dated at 4.9–4.2 Ma, to this suessenbornensis Soergel, 1926 (Rabeder et al., 2010). These form. The exact evo lu tion ary rela tion ships of these bears to au thors also as signed to this form some bears from sites dated Ursavus as well as to later forms re main un clear. In this pa per, to 1.1–0.9 Ma such as Untermaßfeld, Ceyssaguet and var i ous we agree with the as sign ment of these bears to U. minimus as sites in Atapuerca. Pre vi ously, these spec i mens had been de - The his tory of bears (Ursidae, Carnivora, Mammalia) from Silesia (south ern Poland) and the neigh bouring ar eas 881 scribed un der var i ous names – U. rodei Musil, 2001 and form group, U. a. suessenbornensis. This was di agnosed as a U. dolinensis García et Arsuaga, 2001 (García and Arsuaga, me dium-sized brown bear which dif fers from ex tant U. arctos 2001; Musil, 2001). We agree with like Ol ive (2006), Rabeder only by hav ing slightly shorter and wider extrem i ties (Rabeder and Withalm (2006) and Rabeder et al. (2010) that these et al., 2010). Propor tion ally shorter and more robust metapodial names are ju nior synonyms of U. arctos. In contrast to this, bones are a charac ter is tic feature of al most all late Mid dle and some au thors (Baryshnikov, 2007; Argant, 2009) con sid ered Late Pleis to cene brown bears. Nar row ing of metapodial bones that these bears are the old est known repre sen ta tives of oc curred as late as in the lat est part of the Late Pleis to cene and U. deningeri (Wag ner, 2010). From 0.9–0.8 Ma there are also in the Ho lo cene (Rabeder et al., 2010; Wagner, 2010). Re- known some early arctoid-like bears, though showing some mains of this bear spe cies have been noted so far in fauna C of spelaeoid or even thibetanus-like fea tures. For ex am ple, there the Zamkowa Dolna cave in Silesia. New exca va tions at this is ma te rial from the Sackdilling cave in Ger many and Žírany in site may pro vide new ma te rial be long ing to this in ter est ing form. Slovakia (Heller, 1956; Ambros et al., 2005; Baryshnikov, 2007; The long pe riod of cold and con ti nen tal cli ma tic con di tions Wag ner and Sabol, 2007; Wag ner, 2010). be tween 480 and 420 ka (MIS 12, Elsterian or San 2 gla cial in The ear li est rep re sen ta tives of spelaeoid bears ap peared in Po land) dras ti cally changed the Silesian and Palaearctic faunal Eu rope ap proxi mately at the same time as did the bears from pat tern (Kahlke et al., 2011; Kahlke, 2014). It cov ered al most all the arctoid lin eage, and the pe riod be tween 1.4 and 1.2 Ma is of Po land and also al most the entire Silesian terri tory. Spe cies re garded as a splitting time of both lin eages (Wag ner, 2006, of Cen tral Asian steppe or i gin spread into north ern and west ern 2010). From the time be tween 1.1 and 0.9 Ma, the records of Palaearctic re gions. This was part of a lon ger pe riod lasted be- early U. deningeri are rare, and known only from a few Eu ro - tween 0.8 and 0.4 Ma, and char ac ter ized by the begin ning of a pean sites such as Vallonet cave (Moullé, 1992; Baryshnikov, new epi sode in Earth his tory, called the Mid-Pleis to cene Tran si - 2007; Argant, 2009; Wagner, 2010) and Les Valerots in France tion or Mid dle Pleis to cene Rev o lu tion (MPR) (Mas lin and (Erbaeva et al., 2001), Cal Guardiola in Spain (Madurell- Ridgwell, 2005; Clarck et al., 2006). Low-am pli tude, 41 ky cli- Malapeira et al., 2009), and Honce (Wag ner and Sabol, 2007) mate cycles were re placed pro gres sively by high-am pli tude, and Mosbach 1 in Ger many (von Koenigswald and Tobien, 100 ky cy cles. The lat ter cy cles im plied a tran si tion to wards a 1987). However, some au thors assigned the ma te rial from the non-lin ear forced cli mate sys tem, and were ac com pa nied by a Vallonet cave to the arctoid lin eage (García, 2003), and its sub stan tial in crease in global ice vol ume at 0.94 Ma (Mas lin conspecificity with U. arctos sensu Rabeder et al. (2010) can not and Ridgwell, 2005). These cli mate changes, par tic u larly the in- be to tally rejected (Wag ner, 2010). creas ing se ver ity and dura tion of cold stages, had a pro found Re cords of U. deningeri dur ing the lat est Early Pleisto cene ef fect on biota and the phys i cal land scape, espe cially in the (1.0–0.8 Ma) are also very sporadic. Among these, there are North ern Hemi sphere (Kahlke et al., 2011; Kahlke, 2014). Ger man sites Dorn-Dürkheim 3 (Franzen, 1999), Hohensülzen One of the very first fau nal as sem blages termed the Mam - (Storch et al., 1973), Würzburg-Schalksberg (Mäuser, 1987), moth Fauna, as so ci ated with steppe-tun dra con di tions, was the and Rus sian lo cal i ties Port-Katon (Bajguševa et al., 2001) and fauna re corded from melt wa ter grav els de pos ited at ~460 ka in Akhalkalaki (Baryshnikov, 2007). Since the Mid dle Pleis to cene, Bad Frankenhausen; this lo cal ity com prises more spe cies this bear be came com mon and abun dant in most of Eu rope and adapted to cold and con ti nental en vi ron ments, such as Bi son bears from the deningeroid and later spelaeoid lin eage be came sp., Soergelia elisabethae, Praeovibos priscus, Rangifer dom i nant un til the end of the Pleis to cene. Re mains of tarandus ssp., Equus sp. and Mammuthus trogontherii (Kahlke U. deningeri were ob tained from the fol low ing early Middle and Lacombat, 2008; Kahlke, 2014). El ements of steppe and Pleis to cene lo cal i ties: Gombasek (Slovakia; Kretzoi, 1938; tun dra or i gin co-oc curred, and the struc ture of the Mam moth Wag ner and Gasparik, 2014); cave C718, Konìprusy caves, Fauna ap peared for the first time. Chlum 1 and 4 (Czech Re pub lic; Fejfar, 1976; Fejfar and Hein - The period be tween 420 and 300 ka was the time of the for - rich, 1983; Wag ner 2004, 2005, 2010); Voigstedt (Ger many; ma tion of a large car ni vore guild which par tially sur vived un til Horáèek and Ložek, 1988), Jagsthausen (Ger many; Koby, the Ho lo cene and dom i nated over the last 300 ky. The stem of 1952), Kövesvárad (Hun gary; Jánossy, 1963), Kozi Grzbiet this paleoguild com prised Canis lupus ssp., Vulpes vulpes, (Po land; Wiszniowska, 1989), Slivia (It aly; Ambrosetti et al., Ursus spelaeus group, Ursus arctos ssp., Gulo gulo, Panthera 1979), and Tiraspol (Moldova; Da vid, 1982). spelaea ssp., and Crocuta crocuta spelaea. Their re mains though are out num bered in fos sil sites by the bones of spelaeoid bears, con sti tut ing up to 98% at some lo cal i ties; this MIDDLE PLEISTOCENE–HOLOCENE bear was not a carni vore, but a her bi vore, and in rare cases an (700 ka – PRESENT) om ni vore (Marciszak et al., 2011, Krajcarz et al., 2016). Ursus ex gr. arctos The main com pet i tor to large ac tive car ni vores such as the steppe wolf Canis lupus spelaeus, the cave lion Panthera Some prob lem atic spec i mens that may also belong to spelaea spelaea and the Crocuta crocuta spelaea, arctoid bears have been found in ma te ri als from the early and was Ursus arctos priscus. Sim i larly to the steppe wolf, the mid-Mid dle Pleis to cene sites, ~0.8–0.6 Ma (Wag ner, 2010). steppe brown bear is not a dif fer ent spe cies but a form, some - Among them, there are bear re mains from Süßenborn thing like an “eco type”, a spe cial ecomorph adapted to open (Soergel, 1926), West Runton (Rabeder et al., 2010), hab i tats (Marciszak et al., 2016, 2017, 2019a, b, 2020). The Kövesvárad (Jánossy, 1963), Cueva Mayor, Hundsheim great size of this bear, com monly docu mented in Late Pleis to - (Rabeder et al., 2010) and Chlum 4 (Wag ner, 2004, 2010). The cene brown bears from many Euro pean sites, may have been old est arctoid bear in Silesia is from the Po³udniowa cave dated an adap ta tion to colder and more barren hab i tats (Erdbrink, to 0.6–0.5 Ma, but it is not the old est Pol ish re cord of an arctoid 1953, 1967; Ehrenberg, 1955; Kurtén, 1956, 1959, 1968; bear. The ma te rial from Kozi Grzbiet is older; a few iso lated Thenius, 1956; Musil, 1964; Ballesio, 1983; Sabol, 2001a, b; teeth found here may repre sent in fact U. arctos. Based on the Baryshnikov and Boeskorov, 2004; Pacher, 2007; Rabeder and de scrip tion and com par i son of ma te rial from Deutsch-Altenburg Frischauf, 2016; Marciszak et al., 2017, 2019a, b). with other Eu ro pean ma te rial, Rabeder et al. (2010) sug gested Among thou sands of bones, from time to time, arctoid bears be tween 1.4 and 0.6 Ma com pose a rather uni - re mains of the arctoid bear, which is dis tinct in size and mor- 882 Adrian Marciszak and Grzegorz Lipecki

Fig. 3. Skulls of differ ent bears from Silesia and neighbour ing areas A – Ursus ingressus (>) from NiedŸwiedzia cave, Po land (50–40 ka, coll. no. ZP UWr/JNK/Us/1); B – Ursus minimus (>) from Wê¿e 1, Po land (3.6–3.2 Ma, coll. no. MF/833/1); C – Ursus arctos priscus (>) from Bohdalec, Czech Re pub lic (22.8–22.2 ka, coll. no. R 1916/824); D – Ursus arctos arctos (>) from nad Zagonem cave (Zielona cave, Tatra Mts), Po land (3–1 ka, coll. no. ISEZ MF/7390); all skulls shown in lat eral view, scale bar 100 mm phol ogy from the av er age mod ern brown bear, and has been (Figs. 3 and 4). However, this pat tern is not con stant in na ture clas si fied un der dif fer ent names: U. ferox, U. horribilis, U. and there are pop u la tions liv ing in rel a tively south ern lo ca tions, anglicus, U. priscus, U. arctoideus etc. (see the synon ymy in whose av er age body size, due to e.g. con stant or pe ri odic avail - Erdbrink, 1953 for de tails). Di men sions of some in di vid u als, abil ity of high-pro tein food, was sig nif i cantly larger than those of com para ble in size with big cave bears, and the sim i lar ity in cra - in di vid u als in other pop u la tions liv ing at sim i lar lat i tudes nial and postcranial and es pecially in the dental ma te rial, (Mattson and Merrill, 2002; Baryshnikov, 2007). Duringits long caused some au thors to sug gest pos si ble hy bridi sa tion be - tem poral range, the body size of this spe cies was sub ject to tween these forms (Ehrenberg, 1938). It was de scribed as very strong fluc tu a tions which were in flu enced by var i ous fac tors, large bear, with a flat fore head, broad teeth and pro por tion ally es pe cially the cli mate and the type of avail able food. ro bust and short metapodials (Fig. 3; Musil, 1964; Rabeder and The pres ence of Ursus arctos priscus in cave sites showed Withalm, 2006; Baryshnikov, 2007; Rabeder et al., 2010; that this an i mal reg ularly visited caves, most prob ably in search Marciszak et al., 2019a). Among the teeth, mor pho log i cally par- of food. It is less prob a ble that this large bear denned or hi ber - tic u larly out stand ing from “typ i cal arctoid mor phol ogy” are M2 nated in the time when they were oc cu pied by spelaeoid bears, and m3, broad ened and enlarged in com par i son with equiv a - and most of re mains found in caves be longed to very large and lent teeth of U. a. arctos (Figs. 3 and 4). Rode (1931, 1935) had ro bust an i mals, most prob a bly males. Ra dio car bon dates ob - al ready noted that the teeth of U. a. priscus are in size and mor - tained for Ursus arctos priscus showed that this form oc curred phol ogy half way between those of arctoid and spelaeoid bears. in Silesia through out the en tire MIS 3 and MIS 2. It was pres ent Klaatsch (1906: 282) noted: “As a par tic u lar an i mal from the dur ing even the coolest phases, e.g. MIS 2 (Lipecki and Wojtal, end of the ice age, the cave bear (Ursus spelaeus) has al most 2015; Ersmark et al., 2019; Marciszak et al., 2019a, b). equal im por tance with mam moth and rein deer. At the same How ever, even among dom i nant large car ni vores, lev els of time, the brown bear (Ursus arctos) and the grey bear (Ursus lo cal com peti tion were so high that it re sulted in a mar ginal role ferox) also lived in Eu rope, and their re mains ap peared al ready or even al most to tal ab sence of one of the com pet i tors. For ex - in the Palaeo lithic layers; the last of these tran scended the fe- am ple, a re vi sion of more than 50 sites in the Sudetes Mts roc ity and size of a cave bear, and was the most dan ger ous en - showed a to tal ab sence of the cave hyena; most prob ably it was emy of diluvial man”. Sim i larly, Beushausen (1906: 535) men - outcompeted by the steppe wolf, par tic u larly common in this tioned the pres ence of an “immense grey bear”, which lived at area (Marciszak et al., 2016, 2020). Cli ma tic con di tions also the same time as the brown and cave bear. had a very strong im pact, as which is doc u mented by depos its U. arctos is gen er ally more car niv orous in open land scapes dated to MIS 5b-5a. Arid and cold gla cial con di tions meant that (Bojarska and Selva, 2012). Brown bear as a spe cies is a clas - the Cen tral Eu ro pean main land housed the “stan dard” her bi - sic ex am ple of Berg man’s rule, where an in crease in body size vore guild of the mam moth steppe with Mammuthus to wards the north in the North ern Hemi sphere is ob served primigenius, Coleodonta antiquitatis, Equus ferus (large form), The his tory of bears (Ursidae, Carnivora, Mammalia) from Silesia (south ern Po land) and the neigh bouring ar eas 883

Fig. 4. Com par i son of M2 of dif fer ent bears from Silesia and neigh bour ing ar eas A – Ursavus brevirhinus from Opole 2 (12–11 Ma, ZP UWr/OP2/2); B – Ursus minimus from Wê¿e 1 (3.6–3.2 Ma, coll. no. MZ/177); C – Ursus minimus from Wê¿e 2 (2.4–2.2 Ma, coll. no. ZP UWr/W2/1/1); D – from Po³udniowa cave (600–500 ka, coll. no. ZP UWr/JP/4/1); E – Ursus deningeri from Draby 3 (440–400 ka, coll. no. ZPAL M.11/1); F – Ursus rossicus from Naciekowa cave (60–40 ka, ZP UWr/JNW/Us/417); G – Ursus spelaeus spelaeus from Biœnik cave, layer 14 (150–130 ka, coll. no. ZP UWr/JB/Us/12879); H – Ursus ingressus from Biœnik cave, layer 14 (150–130 ka, coll. no. ZP UWr/JB/Us/12879); I – Ursus arctos priscus from Po³udniowa cave (600–500 ka, coll. no. ZP UWr/JP/5/1); J – Ursus arctos priscus from Draby 8 (440–400 ka, coll. no. ZP UWr/D8/5/1); K – Ursus arctos priscus from Kraków-Spadzista (28.8–28.1 ka, coll. no. ISEA MF/7127); L – Ursus arctos cf. arctos from Biœnik cave, layer 1 (14.7–14.2 ka, coll. no. ZP UWr/JB/Ua/14); M – Ursus arctos arctos from Biœnik cave, layer 1ab (3.4–3.2 ka, coll. no. ZP UWr/JB/Ua/14); all teeth showed in occlusal view, scale bar 20 mm

Megaloceros giganteus, Ovibos moschatus, Rangifer tarandus, mate and en vi ron men tal con di tions. Warm ing at the be gin ning and Bi son priscus. Thermophilous and for est spe cies, such as of MIS 1 caused ice sheet re treat to the north and much of the Palaeoloxodon antiquus, Alces alces, Cervus elaphus, Equus Eu rope grad u ally trans formed from arid, grassy steppes into ferus (small form), Bos primigenius, Bi son bonasus, and Saiga bo real for ests. Enor mous herds of ungulates such as horses, tatarica dis ap peared or oc curred in par tic u lar, re stricted short mam moths, an te lopes, steppe bi son, rhi noc er oses and many time pe ri ods. The car ni vore guild was also strik ingly differ ent. other an i mals, that were adapted to exist in open grass lands, Only the large steppe wolf and above-men tioned gi gantic dis ap peared as for est be came pre dom i nant over grass lands. steppe brown bear are recorded more com monly, while the The large bear could not ob tain enough food to sup port its mas - cave lion and cave hyena, if pres ent, are found in very low num - sive body, even if it could sup ple ment its diet with plants. bers and den si ties. Pre lim i nary analy sis of the teeth of C. l. The pro cess and mech a nisms of U. a. priscus dis ap pear - spelaeus and par tially of U. a. priscus showed some height ened ance are still not re solved. It is pos si ble that it became ex tinct level of durophagy, and suggested they expanded into the be cause of com pe ti tion with other pred a tors due to their greater niche of the absent cave hyena. Sim i larly, the pres ence of U. a. eco log i cal plas tic ity. A more re li able sce nario was that some priscus with highly worn teeth sug gested they were pri mar ily pop u la tions re treated east- and north wards. The com pact geo - bone- and frozen car cass-eating scav engers. At this time, there graph ical range was split into iso lated pop ula tions which sur - was an atypi cal situ a tion in Sudetenland, where a rel atively nar - vived across Eu rope. Size decrease took place, and U. a. row car ni vore guild (steppe wolf, steppe brown bear, wol ver ine, priscus evolved into a more her biv orous form due to en vi ron - and cave lion) focused on rich her bi vore pickings. The fear - men tal changes (Fig. 6). Left overs blended into an abun dant some and im mense steppe brown bear be came so large that in nom i na tive form which was much more her biv o rous and om niv - the past their remains were mis iden ti fied as cave bear, and it orous, and so well-adapted to for est con di tions. Sur vival of U. a. was rather for mi dable kleptoparasite rather than ac tive hunter priscus until the early part of MIS 1 is con firmed by many find - (Fig. 5; Marciszak et al., 2016; Musil, 2018; Marciszak et al., ings of the great brown bear with par tic u larly large and broad 2019a, b, 2020). teeth, sim i lar to those in U. a. priscus. Such finds are quite com - U. a. priscus was well-adapted for sur vival in the Late Pleis - mon in de pos its of the Late Gla cial and postglacial pe ri ods, in to cene of Eu rope, but its size may have been linked to the cli - Den mark (DegerbÀl, 1933), Ger many (Freudenberg, 1914; 884 Adrian Marciszak and Grzegorz Lipecki

Fig. 5. Conflict scene between female of Coelodonta antiquitatis and male of Ursus arctos priscus in Skarszyn (Trzebnickie Hills) Such in terac tions were however very rare and usu ally ended in a show of strength. Its size, ex tremely thick skin and long, sharp horns caused that the adult woolly rhino was out of reach of al most all Late Pleis tocene car ni vores. On the other hand, the steppe brown bear was most likely a scaven ger and kleptoparasite. Its im mense pos ture and great size were used to intim i date other carni vores and take their car casses rather that hunt down big prey like rhino or bison. Drawing by W. Gornig

Hilzheimer, 1939), Great Britain (Ad ams, 1880; Reynolds, From the turn of the Mid dle Ages and mod ern times, per ma - 1906), Ire land (Ball and Owen, 1850), Po land (Müller, 1872; nent de struc tion of the en vi ron ment and ex ter mi na tion in - Hilzheimer, 1937; Ruprecht, 1965, 1992), the Neth er lands creased, and the 1600–1750’s were a time of pro gres sive de - (Erdbrink, 1953, 1967), Russia (Müller, 1872; Vereshchagin, cline in pop u la tion and pop u la tion den sity of U. a. arctos in 1959; Baryshnikov and Boeskorov, 2004; Baryshnikov, 2007) Silesia un til its dis appear ance in the mid-18th cen tury. Dur ing and Ukraine (Pidoplichko, 1956). The pres ence of these large this pe riod, hunt ing de scrip tions were a spo radic topic in hunt - brown bears dur ing the postglacial pe riod of Eurasia should be ing chron i cles. Re cords of hunt ing by the Schaffgotsch and re garded as a rel ict sur vival (Baryshnikov, 2007). Tschernin fam i lies show a con stant pres ence of the bear un til Over the last sev eral thou sand years, pro gres sive de vel op- the end of the 18th cen tury. At the same time, however, the ment of set tle ments, de for es ta tion and do mes tic an i mal breed - num ber of an i mals killed clearly shows a de cline in this spe cies ing slowly but sys tem at i cally caused a decrease in the den sity (Pax, 1921, 1925, 1937; Jakubiec, 2001). This spe cies later dis - and pop u la tion of U. a. arctos. How ever, given the low popu la - appeared as a breed ing pop ula tion in Silesia, al though mi grat - tion den sity, the ex tent of for est ar eas and a large num ber of ing in di vid uals were re corded un til the end of the 18th cen tury. In moun tain ar eas, diffi cult to ac cess by humans and not suit able ad di tion, since the Silesian Wars this re gion has passed from for cul ti va tion, this spe cies was quite commonly found through - Aus trian rule into Prus sian, which re sulted in fur ther ad verse out Silesia un til the end of the Mid dle Ages (Pax, 1921, 1925, changes to the bear hab i tat; the Sudetes be came its last ref u- 1937, 1955). Ar chae o log i cal re search show that it in hab ited al - gee. Mas sive de for es ta tion of the lower stream for ests most all of Silesia between the 14th and the 16th cen tu ries, and (Sudetes bank for ests) began and re place ment by spruce its pres ence has been noted in 29 ar chae o log i cal Silesian sites monocultures. Sim i larly, in the Silesian Lowlands, pine was (Wyrost, 1994; Marciszak et al., 2020). Old chron i cles also planted in the place of felled de cid u ous and mixed for ests. Set - men tion mul ti ple re cords of this an i mal in the lowlands and tle ments in creased sig nif i cantly which, com bined with the pro- moun tains of this re gion (Jakubiec, 2001). Be sides, ar chae o - gres sive ex pansion of ag ri cul tural crops and breed ing of do - log i cal and palaeontological data sup ported by direct ra dio car - mes tic an i mals, re sulted in a grad ual re duc tion of the space for bon (C14) dat ing doc u ment, the pres ence of U. a. arctos in bears. The Prus sian method of ra tio nal iz ing for est life pro jected Nowy Targ in Wroc³aw and in rock shel ters on Mi³ek Mt to tal in ter fer ence and re con struction of the food chain. In this (Kaczawskie Mts) be tween 1600 and 1650 has been re corded scheme, there was no place for car ni vores, and thus for a (Ap pen dix 2). brown bear. In di vid u als shot in low land ar eas were mi grat ing in - di vid u als. The Sudetes (and moun tains as well) remained the The his tory of bears (Ursidae, Carnivora, Mammalia) from Silesia (south ern Po land) and the neigh bouring ar eas 885

Fig. 6. Mandi bles of arctoid and spelaeoid bears from Silesia and neighbour ing areas A – Ursus spelaeus spelaeus (>) from Biœnik cave, layer 14 (180–130 ka, coll. no. ZP UWr/JB/Us/144); B – Ursus ingressus (>) from Ciemna cave, layer 2 (35–25 ka, coll. no. ISEZ JC/1061); C – Ursus rossicus (>) from Naciekowa cave (60–40 ka, coll. no. JZP UWr/JNW/Us/12); D – Ursus arctos priscus (>) from NiedŸwiedzia cave (>) (60–50 ka, coll. no. ZP UWr/JNK/Ua/1); E – Ursus arctos priscus (>) from Imbramowice (>) (130–115 ka, coll. no. M.1); F – Ursus arctos arctos (>) from Wschodnia cave, Po land (14.3–13.2 ka, coll. no. MMW/I/A/3927); G – Ursus arctos arctos (+) from cave no. 3 on Birów Hill, Po land (4.3–4.1 ka, coll. no. ZP UWr/C3/Ua/2); all man di bles showed in buccal view, scale bar 100 mm place of their last oc cur rence. In Sudetes Mts, the last in di vid ual MIDDLE PLEISTOCENE–HOLOCENE was killed in the mid-19th cen tury. However, the last Sudetes (700 ka – PRESENT) brown bears were only indi vid u als, which came from the Czech Ursus ex gr. deningeri–spelaeus side from Jesenik and from the Carpathians (Jakubiec, 2001; Flousek et al., 2014). th The tax onomy of the Eu ro pean deningeroid bears has been Af ter the fi nal dis ap pear ance of the spe cies in the mid-18 re viewed by Bishop (1982), Kurtén and Poulianos (1977), Wag - cen tury to the mid-19th cen tury, in di vid u als mi grat ing from the ner (2010), Wag ner and Èermak (2012) and oth ers. In the past Carpathians in Jesenik were observed and even hunted most au thors re cog nised a num ber of spe cies or dif fer ent forms (Jakubiec and Buchalczyk, 1987; Jakubiec, 2001; Flousek et that seemed to re flect dis tinct stages in a progres sively evolv ing al., 2014). For the next 140 years there were no docu mented lin eage which is charac ter ized by gradual changes in den tal bear oc cur rence in the Sudety Mts. It is pos si ble that there were mor phol ogy and body size (Partfitt, 1999). Three sub spe cies of some in di vid uals, but the chron i cles are si lent about such U. deningeri are re cog nised: Ursus deningeri savini (An drews, cases. No in ci den tal cap ture, shoot ing or sight ing of the bear 1922) in Britain and Ursus deningeri suevicus Koby, 1952 in was re corded at that time. Con sid er ing the de gree of pen e tra - con ti nen tal Eu rope, from the lat est Early and early Mid dle Pleis - tion and in dus tri al iza tion of the Sudetes, and their use by tour - to cene, and Ursus deningeri deningeri (von Reichenau, 1904) ists, it seems unlikely that any bear would es cape hu man at ten - which existed in the mid and late Mid dle Pleis to cene. Two these tion. It was not un til 1991–1998 when bear traces were found in sub spe cies are sim i lar in den tal mor phol ogy, and U. d. suevicus the Pol ish Sudetes area (Jakubiec, 1995, 1996). When this is con sid ered less evolved than U. d. deningeri. Some of the ob - bear even tu ally left the Sudetes, over the next 20 years sev eral served dif fer ences may have been due to geo graph ical rather un con firmed ob ser va tions of this spe cies were made, near than to tem po ral and/or phyletic vari abil ity. places such as Wa³brzych, Ludwikowice K³odzkie and The mid-Mid dle Pleis to cene (MIS 16-13) is a typi cal level for G³uszyca (Èervený et al., 2004; Flousek et al., 2014). U. deningeri, with sites like Mosbach 2 (von Reichenau, 1904), 886 Adrian Marciszak and Grzegorz Lipecki

Hundsheim (Zapfe, 1948), Erpfingen 4 (Heller, 1975), La U. ingressus are younger than 37 ka BP (Stiller et al., 2014). Romieu (Prat and Thibault, 1976), Cueva Mayor (García, However, this sce nario of the re place ment of ear lier na tive 2003), Château (Argant and Argant, 2002), West bury (Bishop, spelaeoid forms by U. ingressus can not be re garded as a gen - 1982), Mauer (von Reichenau, 1906), Isernia la Pineta eral ten dency since there are known cases where dif fer ent (Azzaroli et al., 1986), and Petralona (Kurtén and Poulianos, spelaeoid bears co-oc curred over a lon ger pe riod. For exam ple, 1981, Tsoukala, 1991). in two other Austrian caves, Ramesch and Gamssulzen, lo- U. deningeri con tin ued un in ter rupted to the late Mid dle cated ~10 km from each other, U. ingressus lived side by side Pleis to cene and it was char ac ter ized by an increase of body with U. s. eremus for at least 15 ka (Hofreiter et al., 2004). size and cheek-teeth com plexity (Wag ner, 2010). From this pe - Such a re place ment has not yet been recorded in Silesia. riod this spe cies is re corded from Draby 2, 3, and 5, and the More in ter est ing is that U. ingressus co-oc curred with lower most lay ers (19ad-18) of the Biœnik cave dated to MIS U. rossicus in the Kaczawa Mts for a long time, since the youn- 10-8 (Marciszak et al., 2011). Since the bears of spelaeoid lin - gest date for U. rossicus is ~43.5–41.7 ka BP, and it is known eage appeared in layer 15 (MIS 7) and in the younger layers of that both spelaeoid bears were present in the Kaczawa Mts al - the Biœnik cave, this is their old est re cord in the Silesian area ready from at least 80 ky. Dates (ra dio car bon and ura nium-tho- and entire modern ter ri tory of Po land. The evo lu tion of rium) sug gest the more or less con tin u ous pres ence of spelaeoid bears from a deningeroid fore run ner is dif fer ently spelaeoid bears in the Sudetes Mts dur ing the Late Pleis to cene dated at Eu ro pean sites, with the ear li est ap pear ance of (Nadachowski et al., 2008; Wojtal et al., 2015; Marciszak et al., U. spelaeus s. l. noted from the Eng lish site Swanscombe al - 2020). Si mul ta neously, ge netic in ves ti ga tion of in di vid u als from ready during MIS 11 (Kurtén, 1959). However, the ap pear ance Cen tral Eu rope (mainly the Czech Re public, Po land, Slovakia of this spe cies over most of Eu rope took place later, and gen er - and Ukraine) revealed that U. ingressus was the sin gle form of ally it is ac cepted that U. spelaeus s. l. appeared in MIS 7 cave bear pres ent in this part of Eu rope (Popoviæ et al., 2015). (Hilpert, 2006). How ever, it should be noted that ge netic anal ysis of fos sil ma te - The cave bear U. spelaeus s. l. was an ob li gate her bi vore rial from many sites like Biœnik cave failed, and this data has so and one of the most wide spread large mam mals in the Late far not been cor re lated with morphometric anal ysis. By contrast Pleis to cene in Eu rope (Hofreiter et al., 2002; Rabeder et al., with ge netic re sults, morphometric in ves ti ga tions showed the 2004a, b; Baca et al., 2016). The two main Eu ro pean forms, pres ence of at least three dif fer ent cave bear chronosub - U. s. spelaeus and U. ingressus sep a rated prob a bly be tween species, with the dom i nance of U. ingressus since of 420 and 180 ka (Knapp et al., 2009). U. s. spelaeus lived mainly ~100–90 ka BP (Baca et al., 2014). The mtDNA haplotypes of in Western Eu rope, al though it has also been re corded in the in di vid uals from the NiedŸwiedzia cave formed a di ver gent clus - Altai region (Rabeder et al., 2004b; Knapp et al., 2009; Baca et ter on phylogen etic trees, which indi cates the early sepa ra tion al., 2017), while U. ingressus in habited mostly south-east ern and ex pan sion of this pop u la tion (Baca et al., 2012, 2014, and Cen tral Eu rope (Rabeder et al., 2004b; Baca et al., 2014). 2017). Simi lar mtDNA haplotypes have been noted far ther west The geo graphic or i gin of U. ingressus is still not known, but in Zoolithen cave (Stiller et al., 2014). the basal po si tion of haplotypes from the Ro ma nian site The tim ing and causes of the ex tinc tion of spelaeoid bears Peêtera cu Oase in the phylog eny of this spe cies points to SE in Silesia and in Eurasia also re main un clear. Radio car bon Eu rope (Baca et al., 2012). The phylogeographic pic ture of dates show that the last cave bears went extinct prior to the Last U. ingressus is un clear as the mtDNA phylog eny lacks sig nif i - Gla cial Max i mum (LGM). In the past it was thought that they cant sup port and clear phylogeographic lineages can not be dif - dis ap peared ap prox i mately syn chro nously in dif fer ent parts of fer en ti ated. Most prob a bly the spread of U. ingressus may have Eu rope around ~28.5–28 ka at the end of GI-3 (Hofreiter et al., pro ceeded in de pend ently along the main Eu ro pean moun tain 2002; Pacher and Stu art, 2009; Bocherens et al., 2014). ranges, the Alps and the Carpathians (Baca et al., 2014, 2017). Paleogenetic analy ses showed, however, that this pro cess Pre vi ous re search sug gested that be tween 60 and 50 ka started much ear lier, ~50–45 ka (Stiller et al., 2010). U. ingressus started its migra tion west wards along the Alps Stiller et al. (2014) pro posed that spelaeoid bear pop ula - (Hofreiter et al., 2004; Rabeder and Hofreiter, 2004; Münzel et tions might have de clined from east to west, since most of the al., 2011). This opin ion was based on the ear li est and wes tern - sam ples younger than 30 ka have been found in West ern Eu - most records of this spe cies from the Austrian and Swiss Alps, rope. However, AMS dates ob tained over the last few years dated to ~50 ka (Rabeder and Hofreiter, 2004). However, more clearly indi cate that this is not true and that late cave bears sur - re cent data showed that some spelaeoid bears from the Pol ish vived in de pend ently in iso lated pop u la tions in dif fer ent parts of Jura yielded mtDNA haplotypes identi cal to those found in U. Eu rope, even into the mid dle of the GS-3 stadial. In the con text ingressus from the Alps. This may suggest that the pop u la tion of over whelm ing dis ap pear ance, some karst re gions, some - from East ern and SE Eu rope spread north wards be yond the thing akin to a lo cal ref uge, may have pro vided a suit able mi cro - Carpathian Arc (Popoviæ et al., 2015; Baca et al., 2017). cli mate for long sur vival (Baca et al., 2016). An ex am ple of such Ac cord ing to some re search ers, the ar rival of U. ingressus an area in Silesia is Jura Polska, from where the youngest ge - was as so ci ated with the van ish ing of other cave bear forms that net i cally confirmed cave bears so far come from the Stajnia in hab ited the area. One of such re place ments was docu mented cave, dated to ~26.1 ka BP (Baca et al., 2016, 2017). Thus, the from three caves in the Ach Val ley (Ger many; Hofreiter et al., extinc tion time of spelaeoid bears can be es ti mated to be tween 2007; Münzel et al., 2011). The lat est re cords of the na tive in - 27.0 and 24.3 ka BP (Baca et al., 2016, 2017; Mackiewicz et al., habit ant in this area, U. s. spelaeus, were dated to ~31.5 ka BP 2017). while the ear li est ap pear ance of U. ingressus was dated to Al though, in gen eral per cep tion, most of the megafaunal el - 36.3 ka BP. Be side this sin gle old est date, most other speci - ements sur vived un til the end of the Late Pleis to cene, de tailed mens were dated to ~32 ka BP. That sug gests that the main im - study shows that this was not the case. In some ar eas like in mi gra tion of U. ingressus took place just be fore the lo cal extinc - Sudetes Mts this “steppe-tun dra fauna” had van ished al ready tion of U. s. spelaeus. A sim i lar re place ment was doc u mented between 35 and 30 ka, ear lier than in other Pol ish ar eas such as in the Aus trian site Herdengel cave, where the ar rival of the Jura Polska. Cave bears did not sur vive there dur ing the U. ingressus re sulted in the dis appear ance of U. s. eremus. All LGM (Mackiewicz et al., 2017). The van ish ing of these U. s. eremus in di vid uals were dated to >60 ka BP, whereas all megafaunal el ements is only a small part of a gen eral Eu ro pean The his tory of bears (Ursidae, Carnivora, Mammalia) from Silesia (south ern Poland) and the neigh bouring ar eas 887 ten dency, and the extinc tion time co in cides with the end of the were spelaeoid bears, and could win a com pe ti tion for food and first megafaunal tran si tion, which started in Green land hi ber na tion sites, es pe cially during the cold and arid phases. interstadials 5 to 7 (GI-5 to GI-7), and fin ished at the be gin ning Due to its size, com pa ra ble to or even ex ceed ing the size of of the LGM in northern Eu rope (Coo per et al., 2015). There fore, cave bears, it might be re garded as a se ri ous threat to the the ex tinc tion of most mem bers of the so-called “Sudetan spelaeoid bears, and to gether with cave lion and cave wolf as megafauna” repre sents a much ear lier pre-LGM megafaunal the main carni vores feeding on cave bears. This hypoth esis still dis appear ance, a few thou sand years before their fi nal extinc - re quires fur ther test ing by iso to pic stud ies. tion in ar eas like Jura Polska. The decline of the spelaeoid bear Palaeopathological signs on the teeth and bones are found pop u la tions in Eu rope started be tween 50 and 45 ka (Stiller et on many speci mens as signed to dif fer ent spe cies. We sup pose al., 2010), and their fi nal extinc tion was dur ing the cold est and that some bone in ju ries re sulted from trauma bite damage the lon gest (~4 ky) stadial GS-3 (27.5–23.5 ka) in the last gla cial caused by other bears or large car ni vores (mainly their ca nines) (Mackiewicz et al., 2017). from inter- or intraspecies con flicts. The mor tal ity of car ni vores, Cli mate cool ing could have been the main fac tor af fect ing es pe cially siblings and cubs, is strongly influ enced by acci dents cave bear ex tinc tion and re treat of most her bi vores from the like fall ing rocks or be ing stuck in nar row pas sages, ill ness, Sudetes Mts be fore and dur ing the LGM (Stu art and Lis ter, hunger (e.g., in spelaeoid bears in cases when they do not ac - 2007; Pacher and Stu art, 2009; Baca et al., 2016). Spelaeoid cu mu late enough fat dur ing the veg e ta tion sea son), sea sonal bears were strict her bi vores (Kurtén, 1976; Bocherens et al., and ac ci dental floods, which flowed through the cave and 2006; Van Heteren et al., 2009, 2014; Mackiewicz et al., 2010; drowned sleep ing bears, and fi nally simply through old age. Krajcarz et al., 2016). As a con se quence of the cli mate de te ri o - How ever, equally impor tant was the direct ac tiv ity of the main ra tion, the qual ity and availabil ity of plant food in Cen tral Eu - large car ni vores, P. s. spelaea and C. l. spelaeus. Nu mer ous rope, be ing the cru cial com po nent of their diet, had de creased. cave bear bones be longed mostly to cubs and young spec i- The tem per a ture drop could also have af fected the hi ber na tion mens, less than one year old, but also many bones of adult an i - period of spelaeoid bears (Rabeder et al., 2000; Mackiewicz et mals with bite marks, chewing and scratches, are clear in di ca - al., 2017), when this an i mal was more vul ner able to hunt ing tions of pre dation. One of the most spectac ular exam ple is a (Bocherens et al., 2011a, b; Münzel et al., 2011; Diedrich, 2014, neurocranium of the young fe male which holds a few holes, with 2017; Nowakowski and Stefaniak, 2015; Wojtal et al., 2015). two main ones on the fron tal and right pa ri etal bones Among the main fac tors caus ing cave bear ex tinc tion, chang ing (Nowakowski and Stefaniak, 2015). The blurred edges of the cli mate has been pro posed (Baca et al., 2016, 2017). exter nal lamellae and sclerotic lines in di cate that the cave bear Craniodental ad ap ta tions (Kurtén, 1976; Mackiewicz et al., fe male sur vived, how ever prob a bly be cause of ill ness caused 2010; Wiszniowska et al., 2010; van Heteren et al., 2014; by these inju ries it died in the next few months. They were in ter - Pérez-Ramos et al., 2018, 2020) and sta ble iso tope (d13C, d15N) preted as result ing from a very strong bite from the canines of stud ies show that spelaeoid bears were strictly her biv o rous an other car ni vore (Nowakowski and Stefaniak, 2015). It might (Bocherens et al., 1994, 1997; Fernández-Mosquera et al., have been caused by a fight with an other cave bear, as such 2001; Münzel et al., 2011; Krajcarz et al., 2016; Bocherens, be hav iour is com mon in mod ern bears (Erdbrink, 1953). Adult 2019). The cli mate changes which be gan af ter GS-3 caused males kill the cubs or ju ve nile spec i mens usu ally by bit ing their se vere trans for ma tions in plant com mu ni ties around Eu rope skulls (Mathe son, 1942; Vaisfeld and Chestin, 1994). An other (Helmens, 2014). Veg eta tion sea sons shortened and the avail - pos si ble expla nation is an attack by a cave lion, since the hunt - abil ity of high-qual ity plant ma te rial, which seems cru cial for the ing spe ciali sa tion of this spe cies on bears is well-known from ni- sur vival of spelaeoid bears, sig nif i cantly decreased. The di etary tro gen iso tope pat terns (Bocherens et al., 2011a). The skull de - habits of these bears did not fol low any partic ular mod i fi ca tion scribed was found in the NiedŸwiedzia cave, where an al most dur ing their pres ence for the last 10 ky in Eu rope and this eco- com plete skull of spelaea of con sid er able log i cal niche con ser va tism may have led to their de cline size was also found (Wiszniowska, 1978; Barycka, 2008; (Bocherens et al., 2014; Baca et al., 2017). Marciszak et al., 2014). The dis tance be tween two main holes An other crucial factor of extinc tion was the im por tance of (~ 85 mm) fits well with the ca nine spac ing in the lion spec i men. caves for spelaeoid bears as hi ber na tion sites, for which they Ad di tion ally, the cra nium has cut marks, sharp-edged lon gi tu di- had to com pete with hu mans and other car ni vores (Turner, nal scars in di cat ing the use of a sharp-edged tool which was in - 2009; Münzel et al., 2011; Diedrich, 2014; Mackiewicz et al., ter preted as the ef fects of skin ning and pos si ble in di rect proof 2017). Partic ular caves, like NiedŸwiedzia cave, might have of hu man pres ence in the cave (Nowakowski and Stefaniak, been in hab ited by popu la tions where they were born and 2015). However, in our opin ion, an other ex pla nation is also formed sta ble ma ter nal so cial groups for hi ber nation, as was prob a ble. Re cently, when the Si be rian ti ger Panthera tigris found in the case of cave bears from Spain (Fortes et al., 2016). tigris has hunted brown bear, it jumps on a bear from above, The fi del ity to the birth site could de crease the prob a bil ity of and tried to hold its head in its fron tal paws, si mul ta neously bit- find ing a proper hi ber na tion place, in cases where the orig i nal ing through its cer vi cal ver te brae (Ognev, 1935; Mazak, 1979, cave was al ready occu pied or reg u larly vis ited by the main com - 1981; Kirillova and Tesakov, 2008; Kirillova et al., 2009). Dur ing pet i tors/en e mies such as lions, hye nas, wolves, and hu mans. such fights, the ti ger’s claws usually left sharp-edged lon gi tu di - In this con text, is the role of the brown bear is es pe cially inter - nal scars on the bear’s head, the same as are ob served on the est ing, but still not well-stud ied and un der stood, since there are skull of the cave bear fe male from the NiedŸwiedzia cave. It is known re gions where U. arctos expanded its range and re- rea son able to ex trap o late such hunt ing be hav iour to the cave placed spelaeoid bears lead ing to the latter’s ex tinc tion (Kurtén, lion (Kirillova et al., 2009; Diedrich, 2017). 1968; Turner, 2009). Large caves were used by the brown bear In ad di tion to en vi ron men tal and hab i tat changes, some rel a tively rarely, and they usually hiber nated near the en trance other fac tors might have neg a tively influ enced the spelaeoid (Diedrich, 2017). However, U. a. priscus, a gi ant car niv o rous bears. The di rect threat of hunt ing by humans (Münzel et al., form of the brown bear, oc curred (prob ably also co-oc curred) 2011; Wojtal et al., 2015), as well as com pe ti tion from hu mans to gether with U. a. arctos dur ing the Late Pleis to cene for caves as shel ters (Grayson and Delpech, 2003) also had a (Marciszak et al., 2019a, b, 2020). The steppe brown bear was very strong im pact. Mi to chon drial ge nome anal y ses from sev - eco log i cally much better adapted to a con ti nen tal cli mate than eral Span ish cave sites re vealed that spelaeoid bears, in con - 888 Adrian Marciszak and Grzegorz Lipecki trast to U. arctos, ex hib ited strong fi del ity to the caves where cene bears be long to the ge nus Ursus were clas si fied as Ursus they were born. Fac ing grow ing com pe ti tion as a re sult of im - minimus, and they rep re sent one or two mi gra tion events. Ad di - pact with mod ern hu mans and Neanderthals, such hom ing be - tion ally, a sin gle oc cur rence of Agriotherium insigne was re - hav iour might have also con trib uted to their dis appear ance. corded from the Wê¿e 1 site be tween 3.6–3.2 Ma. For this large The ten dency to come back to the same cave for hi ber na tion car niv o rous bear, it was one of the last Eu ro pean oc cur rences. may have prevented the colo nis ation of new areas and made A few Early Pleis to cene bear re cords are rep re sented by a spelaeoid bears more pre dict able for hunters to find (Fortes et prob a ble an ces tor of the arctoid and spelaeoid bear lin eages, al., 2016; Baca et al., 2017). U. etruscus. The old est rep re sen ta tive of Ursus ex gr. arctos so Be sides the well-es tab lished re cords of U. ingressus (Baca far re corded, U. a. suessenbornensis, is known from the latest et al., 2012, 2014, 2017), abun dant ma te rial of dif fer ent forms of Early Pleis to cene (1.2–0.9 Ma) fauna C of Zamkowa Dolna spelaeoid bears was also recov ered from some caves on Mt. cave. A num ber of other Silesian and Pol ish re cords of this form Po³om (Marciszak, 2011). Es pe cially rich bone ma te rial co mes re quire con fir ma tion. The first oc cur rence of U. deningeri, the from the Naciekowa cave, where >7000 bones have been old est taxon of the U. ex gr. deningeri–spelaeus lin eage, was found; in ad di tion, re mains of this spe cies also came from the re corded from the Przymi³owice C site dated to ~700 ka. The re - Wschodnia and prob a bly from the Obok Wschodniej caves. mains of U. deningeri are reg u larly noted from Mid dle Pleis to - The pres ence of an atypi cal pop u la tion from the Wschodnia cene sites within Silesia. cave was al ready recorded by Zotz (1939); he men tioned the Af ter the pro nounced cold pe riod of MIS 12, when the Scan - pres ence of a dwarf race of U. spelaeus with many arctoid mor - di na vian ice sheet cov ered al most all of Po land, ex cept of the pho log i cal fea tures. Zotz (1939) in ter preted the pres ence of this Sudetes and the Carpathians, the fauna changed con sid er ably. form as one of the very last pop u la tions of the cave bear, sim i lar This dras tic fau nal turn over led to the for ma tion of the pan-Eur - to those from the Drachen cave. The ma te rial from the asian Mam moth Fauna at ~460 ka BP. That was a time when Naciekowa cave is char ac ter ized by small in di vid u als. Also the the very char ac ter is tic steppe brown bear Ursus arctos priscus, mor phol ogy, with nar row and sim ply built premol ars and mo lars a spe cial ecomorph adapted to live in open grass lands, ap - (P4 with out ad di tional cusps, m2 and m3 with a poorly devel - peared among the bear fauna. It sur vived un til the be gin ning of oped, straight talonid) and the gracile meta carp als and meta tar - MIS 1, when the mod ern U. arctos arctos appeared in Silesia sals, clearly distin guish the pop u la tion from the Naciekowa and sur vived to the pres ent day. cave from the large and ro bust U. ingressus from the U. deningeri was the most com mon bear in the Mid dle Pleis - NiedŸwiedzia cave (Marciszak, 2011). Mor pho log i cally and in to cene, while the first records of U. spelaeus spelaeus ap- size the small cave bear from the Pro³om cave re sem bles the peared from MIS 7; later, from the Late Pleis to cene small Cau ca sian form called the steppe cave bear U. rossicus (~110–100 ka) it was slowly replaced in Silesia by U. ingressus. (Borissiak, 1930, 1932). Di rect dat ing meth ods (C14 and UTh) Spelaeoid bears to tally domi nated the cave as sem blages, and and iso to pic anal ysis are needed to clar ify its tax o nomic po si - fi nally vanished be tween 27 and 24 ka. The most proba ble fac - tion. tors lead ing these an i mals to their extinc tion were en vi ron men - tal turnover ex ac er bated by clima tic changes and human im - pact. CONCLUSIONS Ac knowl edge ments. We are very grate ful to O. Kovalchuk for lin guis tic improve ment and Alejandro Pérez-Ramos and an - Re vi sion of the Silesian bear fauna based on 155 sites, other anon ymous re viewer for their help ful crit i cal com ments mainly cave and karstic lo cal i ties, but also ar chae o log i cal and and sug ges tions. This work was sup ported by the Na tional Sci- open-air sites, showed the pres ence of 13 bear forms and spe - ence Cen tre of Po land, to Adrian Marciszak un der grant en ti tled cies. These re cords covered the timespan of the last 16.5 Ma; “The his tory of the brown bear Ursus arctos Linnaeus, 1758 they were di vided on five main time groups. The old est bears, evo lu tion in Cen tral Eu rope as a key to mod ern spe cies con ser - rep re sented by the gen era Ballusia and Ursavus, are dated on va tion” no. UMO-2015/17/D/ST10/01907. The re search was 16.5–11 Ma, and be longed to stem forms of the subfamily also sup ported by the Min is try of Sci ence and Higher Ed uca - Ursinae. The old est Silesian bear B. elmenensis was re corded tion, Po land (pro ject 1076/S/IBŒ/2017) and in ter nal grant for from Przeworno 2, dated to 16–14.5 Ma, while the younger Mio - young sci en tists by the Fac ulty of Bi o log i cal Sci ences (pro ject cene bears were assigned to the ge nus Ursavus. no. 0411/2093/18). Af ter a break of 6 My, the ear li est mem bers of the ge nus Ursus ap peared in Pañska Góra at 4.9–4.2 Ma. All known Plio-

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